Life Habits: Benthic



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Zacanthoides romingeri

Zacanthoides romingeri (figure 3) illustrated by Rominger (1887) as Embolimus spinosa.

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Corynexochida
Species name: Zacanthoides romingeri
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Rominger
Description date: 1887
Etymology:

Zacanthoides – probably from the Greek z(a), “very,” and akanthion, “thistle” or “porcupine” or “hedgehog,” and oides, “resembling;” thus, very thistle- or porcupine-like.

romingeri – after Carl Rominger, a Michigan paleontologist who in 1887 published the first descriptions of trilobites from Mount Stephen.

Type Specimens: Type status under review – UMMP 4871 (2 specimens), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Zacanthoides sexdentatus, Z. submuticus, Z. longipygus, Z. planifrons, Z. divergens, all from older and younger Middle Cambrian rocks on Mount Stephen, Mount Odaray, and Park Mountain (Rasetti, 1951).

Other deposits: other species elsewhere in North America.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Trilobite Beds on Mount Stephen.

History of Research:

Brief history of research:

In 1887 Carl Rominger published an engraving of a nearly complete and markedly spiny trilobite and named it Embolimus spinosa. In 1908 Charles Walcott introduced the combination Zacanthoides spinosus for the Mount Stephen species and for a similar trilobite from Nevada. The next change came in 1942, when Charles Resser at the United States National Museum asserted that the Mount Stephen species was sufficiently distinct that it required a new name. Resser chose to honour the man who first formally described many of the common Mount Stephen trilobites, and Zacanthoides romingeri remains the combination in use today.

Description:

Morphology:

Hard parts: adult dorsal exoskeletons can reach up to 6 cm in length, tapering back from a large crescentic cephalon through a thorax of nine segments, to a relatively small rounded-triangular pygidium with long marginal spines.

The wide free cheeks bear strong genal spines; short, thorn-like intragenal spines mark the posterior corners of the fixed cheeks. The glabella is long and narrow, slightly expanded forwards. There are four pairs of lateral glabellar furrows; the anterior two pairs are weaker and angled to the front, the stronger posterior two are angled back. Very long narrow eyes that bow strongly outward are located far back on the cephalon. The occipital ring extends rearward into a strong, broad-based spine. Long, blade-shaped terminal spines on the wide pleurae curve progressively more backwards. A slender needle-like spine arises from the axial ring of the eighth thoracic segment. There are four pygidial axial rings; five pairs of marginal spines, each successively shorter, are directed rearwards and extend beyond the tip of the pygidium.

Unmineralized anatomy: not known.

Abundance:

Zacanthoides romingeri is moderately abundant at the Mount Stephen Trilobite Beds but absent from Fossil Ridge. Complete trilobites with the free cheeks in place are very scarce, and this species is mostly found as disarticulated sclerites. Its distinctive characteristics, however, usually allow even isolated pieces to be readily identified.

Maximum Size:
60 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Zacanthoides romingeri adults very likely walked along the sea bed. The overall spinosity of this species may have served as a deterrent to predators, or possibly helped to break up the visual outline of the animal, making it harder to see on the sea floor (Rudkin, 1996).

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.
  • RESSER, C. E. 1942. Fifth contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 101 (15): 1-58.
  • ROMINGER, C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia, 1887: 12-19.
  • RUDKIN, D. M. 1996. The Trilobite Beds of Mount Stephen, Yoho National Park, p. 59-68. In R. Ludvigsen (ed.), Life in Stone – A Natural History of British Columbia’s Fossils. UBC Press, Vancouver.
  • RUDKIN, D. M. 2009. The Mount Stephen Trilobite Beds, p. 90-102. In J.-B. Caron and D. Rudkin (eds.), A Burgess Shale Primer – History, Geology, and Research Highlights. The Burgess Shale Consortium, Toronto.
  • SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. D. 1888. Cambrian fossils from Mount Stephens, Northwest Territory of Canada. American Journal of Science, Series 3, 36: 163-166.
  • WALCOTT, C. D. 1908. Mount Stephen rocks and fossils. Canadian Alpine Journal, 1:232-248.
Other Links:

http://www.trilobites.info/ordcorynexochida.htm



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Worthenella cambria

Worthenella cambria (USNM 57643) – Holotype, part and counterpart. Left, plate 22 of Walcott (1911), showing a retouched image of the original specimen described (figure 2) together with other “worms.” Right, images of the same specimen. Specimen length = 60 mm. Specimen wet – direct light (left column), dry – polarized light (right column). Walcott Quarry.

© SMITHSONIAN INSTITUTION – NATIONAL MUSEUM OF NATURAL HISTORY. PHOTOS: JEAN-BERNARD CARON

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Megacheira (Hou & Bergström 1997).
Species name: Worthenella cambria
Remarks:

Worthenella has long remained problematic (Briggs & Conway Morris 1986). Legg (2013) proposed that it was a multisegmented “great-appendage” arthropod and formally included it within Megacheira. However, the taxon has remained marginal to recent discussions on early arthropod evolution and phylogenetic analyses, probably because the unique specimen is challenging to interpret and does not provide definitive evidence of its affinity.

Described by: Walcott
Description date: 1911
Etymology:

Worthenella – Possibly after the American palaeontologist Amos Henry Worthen, who died in 1888, just as Walcott’s career was taking off.

cambria – from the Welsh Cambria meaning Wales, in reference to the age of the fossil.

Type Specimens: Holotype –USNM57643 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none

Other deposits: none

Age & Localities:

Age:
Middle Cambrian, Wuliuan stage, Burgess Shale Formation (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Worthenella was first described by Walcott from a single specimen in a 1911 monograph dealing with various Burgess Shale worms. Walcott interpreted this animal as a polychaete annelid (or bristle worm), in the same family as the animal Wiwaxia (which is now interpreted as a stem mollusc). However, This interpretation was questioned (Conway Morris, 1979), and the affinities of Worthenella have remained difficult to establish because this singular fossil is too poorly known (Briggs and Conway Morris, 1986). In light of comparisons with other known Cambrian taxa, Legg (2013) included Worthenella in the class Megacheira, positing the existence of a pair of “great appendages” at the front of the animal.

Description:

Morphology:

The animal is very elongate and tubular, bearing a small head and at least 46 segments of similar size, possibly biramous. Longer appendages are present under the head shield, but their precise nature and arrangement is difficult to determine. The gut is straight and the anus is terminal.

Abundance:

This animal is known from a single specimen.

Maximum Size:
6 cm.

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

This arthropod was likely eating small organic particles or preying on minute prey items on the sea floor. The lack of detailed information on appendage morphology precludes further ecological interpretations.

References:

  • BRIGGS, D. E. G. and CONWAY MORRIS, S. 1986. Problematica from the Middle Cambrian Burgess Shale of British Columbia. In HOFFMAN, A. and NITECKI, M. H. (eds.) Problematic Fossil Taxa (Oxford Monographs on Geology and Geophysics N° 5), Oxford University Press & Clarendon Press, New York, 167–183 pp.
  • CARON, J. B. and JACKSON, D. A. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258, 222–256.
  • HOU, X. G. and BERGSTRÖM, J. 1997. Arthropods of the Lower Cambrian Chengjiang fauna, southwest China. Fossils and Strata, 45, 1–116.
  • LEGG, D. 2013. Multi-segmented arthropods from the Middle Cambrian of British Columbia (Canada). JOURNAL OF PALEONTOLOGY, 87, 493–501.
  • VANNIER, J. and CHEN, J. 2005. Early Cambrian food chain: New evidence from fossil aggregates in the Maotianshan Shale biota, SW China. PALAIOS, 20, 3–26.
  • WALCOTT, C. 1911. Cambrian Geology and Paleontology II. Middle Cambrian annelids. Smithsonian Miscellaneous Collections, 57(5), 109–145.
  • WILLIAMS, M., SIVETER, D. J., POPOV, L. E. and VANNIER, J. M. C. 2007. Biogeography and affinities of the bradoriid arthropods: Cosmopolitan microbenthos of the Cambrian seas. Palaeogeography, Palaeoclimatology, Palaeoecology, 248, 202–232.
Other Links:

None



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Kootenia burgessensis

Kootenia burgessensis (ROM 60761). Disarticulated specimen. Specimen dry – direct light (left) and coated with ammonium chloride sublimate to show details (right). Specimen length = 44 mm. Walcott Quarry.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Corynexochida
Species name: Kootenia burgessensis
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Resser
Description date: 1942
Etymology:

Burgess Shale and vicinity: Kootenia dawsoni. However, see below regarding a possible synonymy with the genus Olenoides.

Other deposits: other species attributed to Kootenia are widespread in the Cambrian of North America, and have been recorded in Greenland, China, Australia, and elsewhere.

Type Specimens: Holotype (K. burgessensis) – USNM65511 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (Resser, 1942); Type status under review – (K. dawsoni), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Kootenia dawsoni; Olenoides serratus. (Species of Kootenia are no longer considered different enough from those in Olenoides to warrant placement in a separate genus, but Kootenia is retained here for ease of reference to historical literature).

Other deposits: other species attributed to Kootenia are widespread in the Cambrian of North America, and have been recorded in Greenland, China, Australia, and elsewhere.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus –Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge, and nearby localities on Mount Field; K. dawsoni is known from the Trilobite Beds and elsewhere on Mount Stephen.

History of Research:

Brief history of research:

Kootenia burgessensis was established by Charles Resser based on material Walcott included in K. dawsoni. Kootenia originally appeared as a subgenus of Bathyuriscus in Walcott’s 1889 paper revising many of Rominger’s Mount Stephen trilobite identifications. Walcott named B. (Kootenia) dawsoni after G. M. Dawson of the Geological Survey of Canada as a replacement for what Rominger had illustrated as Bathyurus (?) in 1887. In 1908, Walcott followed G. F. Matthew (1899) in calling this Dorypyge (Kootenia) dawsoni, but regarded Kootenia as a full genus in 1918. Harry Whittington included Kootenia burgessensis in his 1975 redescription of Burgess Shale appendage-bearing trilobites, illustrating a single specimen showing biramous thoracic limbs on one side. In 1994, Melzak and Westrop pointed out that the diagnostic character of Kootenia (depth of “interpleural” furrows on the pygidium) showed intraspecific variability, and argued that Kootenia—and perhaps other dorypygids trilobites—may have to be subsumed within Olenoides. We maintain Kootenia as a separate genus here pending formal taxonomic clarification.

Description:

Morphology:

Hard parts: adult dorsal exoskeletons may reach 5.5 cm in length and are broadly oval in outline. In most general features, Kootenia burgessensis resembles the co-occurring Olenoides serratus, with a semi-circular cephalon bearing genal spines, a thorax of seven segments, and a semi-circular pygidium. In Kootenia, however, spines on the thoracic pleural tips and shorter and blunter, as are those around the margin of the pygidium; interpleural furrows on the pygidium are absent to very faint.

Unmineralized anatomy: based on evidence from just a few specimens, Kootenia burgessensis, like Olenoides serratus, had a pair of flexible, multi-jointed “antennae” followed by three pairs of biramous limbs on the cephalon. Pairs of similar biramous appendages were attached under each thoracic segment, with a smaller number under the pygidium. No specimens, however, show any evidence of posterior antenna-like cerci as in Olenoides.

Abundance:

Kootenia burgessensis is moderately common in the Walcott Quarry section on Fossil Ridge, as is Kootenia dawsoni in the Mount Stephen Trilobite Beds.

Maximum Size:
55 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Adult Kootenia burgessensis walked along the sea bed, possibly digging shallow furrows to locate small soft-bodied and weakly-shelled animals or carcasses. Kootenia could probably swim just above the sea bed for short distances. Tiny larvae and early juveniles probably swam and drifted in the water column.

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • MATTHEW, G. F. 1899. Studies on Cambrian faunas, No. 3. Upper Cambrian Fauna of Mount Stephen, British Columbia: The trilobites and worms. Transactions of the Royal Society of Canada, Series 2, Vol. 5, Section IV:39-66.
  • MELZAK, A. AND S. R. WESTROP. 1994. Mid-Cambrian (Marjuman) trilobites from the Pika Formation, southern Canadian Rocky Mountains, Alberta. Canadian Journal of Earth Sciences, 31:969-985.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.
  • RESSER, C. E. 1942. Fifth contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 101 (15): 1-58.
  • RESSER, C. E. 1942. Fifth contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 101 (15): 1-58.
  • ROMINGER, C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia, 1887: 12-19.
  • SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. 1889. Description of new genera and species of fossils from the Middle Cambrian. United States National Museum, Proceedings for 1888:441-446.
  • WALCOTT, C. D. 1908. Mount Stephen rocks and fossils. Canadian Alpine Journal, 1: 232-248.
  • WALCOTT, C. 1918. Cambrian Geology and Paleontology IV. Appendages of trilobites. Smithsonian Miscellaneous Collections, 67(4): 115-216.
  • WHITTINGTON, H. B. 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia. Fossils and Strata, No. 4: 97-136.
Other Links:

http://www.trilobites.info/ordcorynexochida.htm



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Hanburia gloriosa

Hanburia gloriosa (ROM 48468). Complete individual (external mold). Specimen length = 26 mm Specimen coated with ammonium chloride sublimate to show details. Trilobite Beds on Mount Stephen.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Corynexochida?
Species name: Hanburia gloriosa
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

 

Already translated – see Bathyuriscus

Described by: Walcott
Description date: 1916
Etymology:

Hanburia – unspecified, but probably after Hanbury Peak or Hanbury Glacier in the Canadian Rockies, in turn named for David T. Hanbury (1864-1910), a British explorer of the Canadian Northwest Territories.

gloriosa – from the Latin gloriosus, meaning “glorious” or “boastful,” perhaps in allusion to the unusual cephalic morphology of this rare species.

Type Specimens: Lectotype –USNM61724, in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

Age:
Middle Cambrian, Glossopleura to Bathyuriscus-Elrathina Zones (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge. The Tulip Beds (S7) and smaller localities on Mount Stephen.

History of Research:

Brief history of research:

Walcott’s three original specimens of Hanburia gloriosa were found over the course of five years of quarrying the Phyllopod Bed on Fossil Ridge (Walcott, 1916); two more from this locality are also in theUSNMcollections. A singleUSNMspecimen was later found by Charles Resser, supposedly from the “Ogygopsis shale” on Mount Stephen (Rasetti, 1951), but this is almost certainly an error. Harry Whittington reassessed this odd trilobite in 1998.

Description:

Morphology:

Hard parts: the few known specimens of Hanburia gloriosa range in length from 4 mm (for a juvenile stage) to 35 mm. Dorsal shields are broadly ovate to subcircular in outline and all specimens are considerably flattened by compression of the thin exoskeleton. The cephalon is semicircular with a weak, shallow border furrow along the posterior and lateral margins, fading out towards the anterior corners of the glabella. The glabella in small specimens expands forwards and shows two pairs of faint bulbous lateral lobes; in larger specimens, the glabella is parallel-sided and the lobes are subdued. There are no apparent eyes located laterally on the cephalon, and there is no sign of dorsal facial suture. In these two features, Hanburia is unique among the non-agnostoid trilobites of the Burgess Shale.

Whittington (1998) has suggested that the facial suture might run along the outside edge of the cephalon, or ventrally, crossing to the dorsal side only at the genal angles, which in all specimens appear to be rounded. Larger individuals show six or seven segments in the comparatively short thorax, and a single known (presumed) juvenile stage shows four; the distal tips of the pleurae are rounded. The semicircular pygidium lacks a defined border, and is approximately the same width and length as the cephalon. Seven or eight axial rings and a terminal piece make up the pygidial axis, which ends short of the posterior margin. Eight or nine pairs of well-marked pygidial pleurae radiate out and back from the axis.

Unmineralized anatomy: not known

Abundance:

Very rare in all the Burgess Shale localities.

Maximum Size:
35 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Due to its unusual cephalic morphology (i.e., no dorsal sutures or lateral compound eyes), rarity, and unique occurrence only in the Burgess Shale, Hanburia gloriosa remains an ecological enigma. Other “blind” Cambrian trilobites with somewhat similar morphologies have been interpreted as inhabiting deeper waters, perhaps below the photic zone (Whittington, 1998).

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.
  • SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. D. 1916. Smithsonian Miscellaneous Collections, 64(3): 157-258.
  • WHITTINGTON, H. B. 1998. Hanburia gloriosa: rare trilobite from the Middle Cambrian, Stephen Formation, British Columbia, Canada. Journal of Paleontology, 72: 673-677.
Other Links:

None



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Elrathina cordillerae

Elrathina cordillerae (ROM 53273). Complete individual; a presumed carcass with free cheeks in place (coated with ammonium chloride sublimate to show details). Specimen length = 24 mm. Specimen dry – direct light. Mount Stephen Trilobite Beds on Mount Stephen.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Ptychopariida
Species name: Elrathina cordillerae
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Rominger
Description date: 1887
Etymology:

Elrathina – unspecified.

cordillerae – in reference to the Western Cordillera (Canadian Rocky Mountain ranges), derived from the Spanish cordilla, the diminutive of cuerda, meaning “cord.”

Type Specimens: Type status under review – UMMP 4883 (6 specimens), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Elrathina parallela, E. brevifrons, E. spinifera, and E. marginalis have been described from similar stratigraphic horizons at nearby sites on Mount Field, Mount Stephen, and Mount Odaray.

Other deposits: Other species of Elrathina have been reported from the Cambrian of North America and Greenland.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus–Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge. The Trilobite Beds and additional localities on Mount Stephen.

History of Research:

Brief history of research:

E. cordillerae was originally described under the genus name Conocephalites in Rominger’s 1887 publication on trilobites from Mount Stephen. In 1888 Walcott reallocated the species to Ptychoparia where it remained until Charles Resser, Walcott’s former assistant at the United States National Museum, established the new replacement genus Elrathina (Resser, 1937). Other workers have subsequently suggested that Elrathina is indistinguishable from Ptychoparella (see Blaker and Peel, 1997).

Species of Elrathina, along with those of the corynexochid Bathyuriscus, were found to be very abundant in a narrow interval of Middle Cambrian rocks throughout western North America, forming the basis of the Bathyuriscus-Elrathina Zone erected by Charles Deiss (1940).

Description:

Morphology:

Hard parts: adult dorsal exoskeletons average about 2 cm long. The semicircular cephalon is about one-third the length of the entire dorsal shield, bordered by a well-defined narrow rim, and with rounded genal angles. Weak transverse eye ridges extend to the small eyes, which are located just forward of cephalic mid-length. The slightly anteriorly narrowing glabella is rounded in front and exhibits three pairs of shallow lateral furrows; the pre-glabellar field is about the same width as the narrow anterior rim. The long, tapering thorax with a narrow axial lobe contains between 17 and 19 straight-sided segments, flexed gently downwards a short distance from the rounded tips. The tiny elliptical pygidium usually features two segments.

Unmineralized anatomy: rare specimens from the Walcott Quarry on Fossil Ridge retain tantalizing evidence of soft parts, including a pair of slender uniramous antennae, followed by very delicate looking biramous limbs beneath the cephalon, thorax and pygidium. These and other individuals of E. cordillerae are occasionally associated with a dark stain adjacent to the exoskeleton, presumably representing fluidized decay products.

Abundance:

Relatively common on Fossil Ridge and locally very abundant in the Walcott Quarry, where it represents about 25% of all trilobites collected (Caron and Jackson, 2008).

Maximum Size:
28 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Like similar-looking ptychoparioid trilobites, E. cordillerae may be interpreted as a fully mobile, epibenthic deposit (particle) feeder adapted to very low oxygen levels.

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • BLAKER, M. R. AND J. S. PEEL. 1997. Lower Cambrian trilobites from North Greenland. Meddeleser om Grønland, Geoscience, 35, 145 p.
  • CARON, J.-B. AND D. A. JACKSON. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • DEISS, C. 1940. Lower and Middle Cambrian stratigraphy of southwestern Alberta and southeastern British Columbia. Bulletin of the Geological Society of America, 51: 731-794.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 277 p.
  • RESSER, C. E. 1937. Third contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 95(22): 29 p.
  • ROMINGER, C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia, 1887: 12-19.
  • RUDKIN, D. M. 1989. Trilobites with appendages from the Middle Cambrian Stephen Formation of British Columbia. 28th International Geological Congress, Washington, D.C. July 9-19, 1989. Abstracts: 2-729.
  • SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. 1918. Cambrian Geology and Paleontology IV. Appendages of trilobites. Smithsonian Miscellaneous Collections, 67(4): 115-216.
  • WALCOTT, C. D. 1924. Cambrian and Lower Ozarkian trilobites. Smithsonian Miscellaneous Collections, 75(2): 53-60.
Other Links:

http://www.trilobites.info/ordptychopariida.htm

http://www.geological-supplies.com/burgess/abstracts/balint

http://gsa.confex.com/gsa/2008AM/finalprogram/abstract_149017.htm



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Elrathia permulta

Elrathia permulta (ROM 60762). Complete individual; a presumed carcass with free cheeks in place. Specimen length = 27 mm. Specimen dry – direct light (left) and coated with ammonium chloride sublimate to show details (right). Walcott Quarry.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Ptychopariida
Species name: Elrathia permulta
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Walcott
Description date: 1918
Etymology:

Elrathia – unspecified.

permulta – from the Latin per, “very much”, and multus, “many”.

Type Specimens: Holotype –USNM65517 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: other species occur, sometimes abundantly, elsewhere in the Cambrian of North America and Greenland.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus–Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge. Elrathia sp. has been reported from localities on Mount Stephen.

History of Research:

Brief history of research:

The concept of Elrathia permulta is quite confused. Walcott named the species Ptychoparia permulta in 1918 and illustrated two specimens, one clearly designated as the type. Resser (1937) noted that the illustrated specimens were quite different, moved both to Walcott’s 1924 genus Elrathia, and proposed the name Elrathia dubia for the second species. Unfortunately, he based this on the original type of permulta; Rasetti (1951) declared dubia invalid, returned the type specimen to Elrathia permulta, and designated the other of Walcott’s specimens as a paratype of Ehmaniella burgessensis. The holotype of permulta, however, lacks many of the diagnostic characters of Elrathia, and Robison (1964) suggested it represents a new genus.

Description:

Morphology:

Hard parts: the adult dorsal exoskeleton is up to 25 mm long, with a large semicircular cephalon occupying about one-third the total length. The cephalon is bordered by a rounded rim and broad inner furrow; genal angles are produced into sharp triangular spines extending back to the fourth thoracic segment. There is a relatively long field between the narrow, tapered, and anteriorly rounded glabella and the frontal rim. Eyes are small and transverse eye ridges are very weak. Three pairs of shallow lateral furrows mark the glabella. The thorax comprises 14 segments, and tapers back more rapidly over the posterior half to a small rounded pygidium. The surface of the exoskeleton is variably granulate.

Unmineralized anatomy: not known.

Abundance:

Rare in the Walcott Quarry on Fossil Ridge, and elsewhere.

Maximum Size:
25 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

E. permulta may, like similar small ptychoparioid trilobites, be interpreted as a mobile, epibenthic deposit (particle) feeder adapted to low oxygen levels.

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 277 p.
  • RESSER, C. E. 1937. Third contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 95(22): 29 p.
  • ROBISON, R. A. 1964. Late Middle Cambrian faunas from western Utah. Journal of Paleontology, 38:510-566.
  • SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. 1918. Cambrian Geology and Paleontology IV. Appendages of trilobites. Smithsonian Miscellaneous Collections, 67(4): 115-216.
  • WALCOTT, C. D. 1924. Cambrian and Lower Ozarkian trilobites. Smithsonian Miscellaneous Collections, 75(2): 53-60.
Other Links:

http://www.trilobites.info/ordptychopariida.htm



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Ehmaniella burgessensis

Ehmaniella burgessensis (ROM 60759) – Part and counterpart. Complete specimen. Specimen length = 6 mm. Specimen dry – direct light (left) and coated with ammonium chloride sublimate to show details (middle, right). Walcott Quarry

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Ptychopariida
Species name: Ehmaniella burgessensis
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Rasetti
Description date: 1951
Etymology:

Ehmaniella – modification of Ehmania, a trilobite genus name coined in 1935 by C. E. Resser to honour Philip Ehman (Montana) for his geological assistance.

burgessensis – from the Burgess Shale.

Type Specimens: Holotype (E. burgessensis) – USNM116245; Holotype (E. waptaensis) – USNM116243 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: Ehmaniella waptaensis Rasetti, 1951.

Other deposits: other species have been reported from elsewhere in the Cambrian of North America.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus–Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott, Raymond and Collins Quarries on Fossil Ridge. The Trilobite Beds on Mount Stephen, and smaller localities on Mount Odaray.

History of Research:

Brief history of research:

Walcott illustrated two Burgess Shale trilobite specimens in establishing Ptychoparia permulta in 1918. Resser (1937) saw that the two individuals belonged in different species, but erroneously used Walcott’s clearly designated primary type of permulta to found the new combination Elrathia dubia. Rasetti (1951) declared Resser’s dubia invalid, left the original type of permulta in Elrathia, and employed Walcott’s other specimen as a paratype of a new species (burgessesnsis), which he assigned to Resser’s 1937 genus Ehmaniella. Ehmaniella waptaensis, also described by Rasetti in 1951, is nearly indistinguishable.

Description:

Morphology:

Hard parts: adult dorsal exoskeletons may reach 2.8 cm long. The semicircular cephalon is about one-third the length of the dorsal shield, bordered by a well-defined rounded rim; wide free cheeks often show anastomosing ridges and carry short, sharp genal spines. Strong transverse eye ridges extend to relatively large eyes, which are located at or behind cephalic mid-length. The bluntly rounded glabella tapers evenly forward and bears three pairs of shallow lateral furrows; the pre-glabellar field is short. A thorax of thirteen parallel-sided segments has a barrel-shaped outline and a rather broad axial lobe. The short, wide, rounded triangular pygidium usually shows 4 or 5 axial rings with corresponding pleurae. The surface of the exoskeleton is variably granulate.

Unmineralized anatomy: rare specimens of Ehmaniella from the Walcott Quarry and above on Fossil Ridge preserve a pair of slender uniramous antennae (Walcott, 1918; Rudkin 1989). These are sometimes associated with a dark stain adjacent to the exoskeleton, presumably representing fluidized decay products.

Abundance:

Relatively common on Fossil Ridge and locally abundant in the Walcott Quarry (fourth most common trilobite with about 400 specimens observed, only 13 of which are E. waptaensis, Caron and Jackson, 2008).

Maximum Size:
28 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Like similar-looking ptychoparioid trilobites, Ehmaniella may be interpreted as a fully mobile, epibenthic deposit (particle) feeder.

References:

  • Aria, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • Aria, C., Caron, J.-B. & Gaines, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • Cotton, T.J. & Braddy, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • Paterson, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • Paterson, J.R., Edgecombe, G.D. & Lee, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
Other Links:

http://www.trilobites.info/ordptychopariida.htm



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Olenoides serratus

3D animation of Olenoides serratus.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Corynexochida
Species name: Olenoides serratus
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Rominger
Description date: 1887
Etymology:

Olenoides – from Olenus, in Greek mythology a man who, along with his wife Lethaea, was turned to stone. Olenus was used for a trilobite genus name in 1827; the suffix –oides(“resembling”) was added later.

serratus – from the Latin serratus, “saw-shaped,” probably referring to the spinose margin of the pygidium.

Type Specimens: Type status under review – UMMP 4882 (11 specimens), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Kootenia dawsoni; Kootenia burgessensis. (Species of Kooteniaare no longer considered different enough from those in Olenoides to warrant placement in a separate genus, but Kootenia is retained here for ease of reference to historical literature).

Other deposits: species of Olenoides are widespread in the Cambrian of North America and Greenland, and have been recorded in Siberia, China, and elsewhere.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge. The Trilobite Beds and other localities on Mount Stephen.

History of Research:

Brief history of research:

Olenoides serratus was among the first Burgess Shale animals to be named and described. The fossils used in Rominger’s original 1887 description were collected from the Mount Stephen Trilobite Beds in 1886. Rominger coined the name Ogygia serrata for this trilobite, illustrating one complete specimen in accompanying engravings. Following several intermediate changes, the name now in use was first published by Kobayashi in 1935. Spectacular appendage-bearing specimens discovered during Walcott’s Fossil Ridge excavations in 1910-1911 brought Olenoides serratus (then called Neolenus serratus) attention worldwide as one of the most anatomically complete trilobites known. This iconic Burgess Shale species has been thoroughly redescribed by Harry Whittington (1975, 1980), who also concluded that Nathorstia transitans (named by Walcott in 1912) was a “soft shell” moult stage of Olenoides serratus.

Description:

Morphology:

Hard parts: adult dorsal exoskeletons may reach 9 cm long and are broadly oval in outline, with a semi-circular cephalon, a thorax of seven segments ending in spines, and a semi-circular pygidium with marginal spines. The cephalon, thorax and pygidium are of approximately equal length. The parallel-sided glabella is rounded in front and reaches almost to the anterior border. Thin eye ridges swing back from the front of the glabella to the small, outwardly-bowed eyes. The free cheeks narrow back into straight, slender genal spines reaching to the third pleurae. Tips of the pleurae also extend into needle-like spines. The spiny pygidium has six axial rings decreasing in size backwards; five pairs of marginal spines point rearward. The whole exoskeleton has a variably granulate outer surface with fine ridges and cusps near the margins.

Unmineralized anatomy: Olenoides serratus had a pair of flexible, multi-jointed cephalic “antennae.” Behind these, three pairs of biramous limbs were attached beneath the cephalon on either side of the mid-line. Each inner branch had a large spiny blade-shaped coxa and six spinose cylindrical podomeres that tapered away from the body, the last carrying three short “claws” at the tip. The outer limb branch was composed of many flat, overlapping filaments sweeping back from a long lobe, with a small oval, hair-fringed lobe at the outer end. Pairs of similar biramous appendages were attached under each thoracic segments; four to six pairs were attached under the pygidium, becoming shorter and more slender to the rear. Unique among all trilobites preserving limbs, Olenoides serratus also had a pair of antenna-like appendages (cerci; singular = cercus) emerging from under the pygidium behind the last biramous limbs.

Abundance:

Olenoides serratus is moderately common, especially at the Mount Stephen Trilobite Beds, where thousands of pieces and hundreds of partial to complete exoskeletons have been observed or collected. Olenoides is the largest and most conspicuous trilobite in the Walcott Quarry section on Fossil Ridge, where specimens with preserved appendages have been found.

Maximum Size:
90 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Adults of Olenoides serratus walked along the sea bed, possibly digging shallow furrows to locate small soft-bodied and weakly-shelled animals or carcasses. Prey items were shredded between the spiny limb bases and passed forward to the rear-facing mouth. Olenoides could probably swim just above sea bed for short distances. Some Olenoides fossils show unmistakable evidence of healed injuries, suggesting they may have been preyed upon, likely in their “soft-shell” growth phase, by larger arthropods such as Anomalocaris. Tiny larvae and early juveniles of Olenoides probably swam and drifted in the water column above the sea bed.

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • KOBAYASHI, T. 1935. The Cambro-Ordovician formations and faunas of south Chosen. Paleontology, Part 3: Cambrian faunas of south Chosen with a special study on the Cambrian trilobite genera and families. Journal of the Faculty of Science, Imperial University of Tokyo, Section II. 4(2): 49-344.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.
  • ROMINGER, C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia, 1887: 12-19.
  • SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. D. 1912. Cambrian Geology and Paleontology, II. No. 6. – Middle Cambrian Branchiopoda, Malacostraca, Trilobita, and Merostomata. Smithsonian Miscellaneous Collections, 57(6): 145-228.
  • WHITTINGTON, H. B. 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia. Fossils and Strata, No. 4: 97-136.
  • WHITTINGTON, H. B. 1980. Exoskeleton, moult stage, appendage morphology, and habits of the Middle Cambrian trilobite Olenoides serratus. Palaeontology, 23: 171-204.
Other Links:

http://www.trilobites.info/ordcorynexochida.htm

http://www.trilobites.info/trilovent.htm

http://paleobiology.si.edu/burgess/olenoides.html

http://pakozoic.deviantart.com/art/Olenoides-serratus-3D-77550691



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Bathyuriscus rotundatus

Bathyuriscus rotundatus (USNM 116232b) – Plesiotype. Nearly complete individual with right free cheek in place. Specimen length = 14 mm. Specimen dry – direct light. Trilobite Beds on Mount Stephen.

© Smithsonian Institution – National Museum of Natural History. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Benthic
Phylum: Benthic
Higher Taxonomic assignment: Class Trilobita, Order Corynexochida
Species name: Bathyuriscus rotundatus
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Rominger
Description date: 1887
Etymology:

Bathyuriscus – a variation of the earlier trilobite genus name Bathyurus, originally based on the Greek bathys, “deep,” and the Greek oura, “tail,” thus, a trilobite with a deep tail.

rotundatus – from the Latin rotundus, “round,” presumably alluding to the rounded outline of the dorsal shield.

Type Specimens: Type status under review – UMMP 4884 (9 specimens), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Bathyuriscus adaeus Walcott, 1916, from several localities higher in the Bathyuriscus-Elrathina Zone on Mount Stephen, Mount Odaray, and Park Mountain.

Other deposits: other species of Bathyuriscus have been described from numerous localities elsewhere in the Cambrian of North America.

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus–Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Trilobite Beds and other localities on Mount Stephen. Fossil Ridge in sections stratigraphically below the Walcott Quarry.

History of Research:

Brief history of research:

Bathyuriscus rotundatus was first described in the same 1887 publication as several other important Mount Stephen trilobites. Carl Rominger initially used the name Embolimus rotundata for partial specimens of this trilobite, and named a second similar species in his collection Embolimus spinosa (now known as Zacanthoides romingeri). In 1908, Walcott revised Rominger’s original species name to yield the combination Bathyuriscus rotundatus, still in use today (Walcott, 1908). Along with the co-occurring Elrathina cordillerae, B. rotundatus is a signature fossil for the Middle Cambrian Bathyuriscus-Elrathina Zone in the southern Canadian Rockies.

Description:

Morphology:

Hard parts: adult dorsal exoskeletons may be up to 5 cm long and are narrowly oval in outline, with a semicircular cephalon, a thorax of nine segments ending in blade-like tips with short spines, and a semicircular pygidium without spines. The long glabella reaches almost to the anterior cephalic border; the posterior portion is narrow and parallel-sided, while the anterior third expands rapidly forward. There are four pairs of lateral glabellar furrows, with the two front pairs angled forward and the posterior pair directed obliquely back. The eyes are relatively long and lie close to the glabella. Broad free cheeks are extended back into short genal spines. The pygidium is slightly smaller than the cephalon, with a well-defined narrow axial lobe of five rings and a terminal piece; four pairs of pygidial ribs are usually visible. The exoskeleton is mostly smooth externally, but very well-preserved specimens may show faint anastomosing ridges on the free cheeks. Unmineralized anatomy: not known.

Abundance:

Extremely common in the Mount Stephen Trilobite Beds, where it rivals Ogygopsis klotzi in abundance.

Maximum Size:
50 mm

Ecology:

Life habits: Benthic
Feeding strategies: Benthic
Ecological Interpretations:

Bathyuriscus rotundatus was a mobile epibenthic trilobite. Because we have no direct evidence of limb structure, its feeding habits are uncertain. It may have been a deposit feeder and opportunistic scavenger. Like Ogygopsis, Bathyuriscus may occur as fully intact individuals (probably carcasses), with the free cheeks missing, inverted, or rotated (presumed moults), and as scattered pieces. Some show evidence of healed injuries that may be predation scars (Rudkin, 2009).

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.
  • ROMINGER, C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia, 1887: 12-19.
  • RUDKIN, D. M. 2009. The Mount Stephen Trilobite Beds, pp. 90-102. In J.-B. Caron and D. Rudkin (eds.), A Burgess Shale Primer – History, Geology, and Research Highlights. The Burgess Shale Consortium, Toronto.
  • SCHOLTZ, G. AND EDGECOMBE, G. D. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.
  • WALCOTT, C. D. 1888. Cambrian fossils from Mount Stephens, Northwest Territory of Canada. American Journal of Science, series 3: 163-166.
  • WALCOTT, C. D. 1908. Mount Stephen rocks and fossils. Canadian Alpine Journal, 1: 232-248.
  • WALCOTT, C. D. 1916. Cambrian Geology and Paleontology III. Cambrian Trilobites. Smithsonian Miscellaneous Collections, 64(5): 303-456.
Other Links:

http://www.trilobites.info/ordcorynexochida.htm